Late Cenomanian–Turonian (Cretaceous) ammonites from Wadi Qena, central Eastern Desert, Egypt:...

INTRODUCTION

Marine lower Upper Cretaceous successions arewell exposed in the northeastern part of Egypt (Sinaiand Eastern Desert; Text-fig. 1). The southernmostexposures of Upper Cenomanian–Turonian sediments

are preserved in the central part of the Eastern Desertin Wadi Qena, and ammonites are among the most im-portant faunal elements in these strata. However, onlythree papers from the last century have dealt with thistopic (Eck 1914; Douvillé 1928; Luger and Gröschke1989). Therefore, this paper aims to enhance the

Late Cenomanian–Turonian (Cretaceous) ammonites fromWadi Qena, central Eastern Desert, Egypt: taxonomy, bio-

stratigraphy and palaeobiogeographic implications

EMAD NAGM1 AND MARKUS WILMSEN2

1Geology Department, Faculty of Science, Al-Azhar University, Assiut, Egypt. E-mail: [email protected]

2Senckenberg Naturhistorische Sammlungen Dresden, Museum für Mineralogie und Geologie, Sektion Paläozoologie, Königsbrücker Landstr. 159, D-01109 Dresden, Germany. `

E-mail: [email protected]

ABSTRACT:

Nagm, E. and Wilmsen, M. 2012. Late Cenomanian–Turonian (Cretaceous) ammonites from Wadi Qena, centralEastern Desert, Egypt: taxonomy, biostratigraphy and palaeobiogeographic implications. Acta Geologica Polonica,62 (1), 63–89. Warszawa.

In Egypt, marine Upper Cenomanian–Turonian strata are well exposed in the Eastern Desert. The southernmostoutcrops are located in the central part of Wadi Qena, where the lower Upper Cretaceous is represented by the fos-siliferous Galala and Umm Omeiyid formations. From these strata, numerous ammonites have been collectedbed-by-bed and 13 taxa have been identified, which are systematically described herein. Four of them (Euom-phaloceras costatum, Vascoceras globosum globosum, Thomasites gongilensis and Pseudotissotia nigeriensis) arerecorded from Egypt for the first time. The ammonite ranges are used for a biostratigraphic zonation of the lowerUpper Cretaceous succession in the northern and central part of Wadi Qena: the Upper Cenomanian–Lower Tur-onian has been subdivided into five biozones (including a new upper Lower Turonian biozone based on the oc-currence of Pseudotissotia nigeriensis), and one biozone has been recognized in the Upper Turonian.Palaeobiogeographically, the ammonite assemblage has a Tethyan character. During the Early Turonian, influencesof the Vascoceratid Province were predominant with strong affinities to typical Nigerian faunas. This shows thesignificance of faunal exchange between Egypt and Central and West Africa via the Trans-Saharan Seaway. Com-pared to contemporaneous ammonoid faunas from the northern part of the Eastern Desert, Boreal influences aremuch less obvious in Wadi Qena. Thus, the present study greatly enhances the knowledge of the Late Cretaceouspalaeobiogeography and biostratigraphy of Egypt and adjacent areas.

Key words: Upper Cretaceous; Ammonites; Systematic palaeontology; Biostratigraphy;Palaeobiogeography; Egypt.

Acta Geologica Polonica, Vol. 62 (2012), No. 1, pp. 63–89

64EMAD NAGM AND MARKUS WILMSEN

knowledge of the taxonomy of the Late Cenoman-ian–Turonian ammonite fauna in Wadi Qena by meansof a modern systematic approach and description of alarge fauna of ammonites collected bed-by-bed fromthe Galala and Umm Omeiyid formations. In total, 90ammonites have been collected during two field cam-

paigns (April and June 2011) from two sections inWadi Qena, forming the basis of the study. In addition,a detailed biostratigraphic framework is proposed forthe succession based on the ranges of the ammonites.Finally, the palaeobiogeographic affinities of the am-monite assemblage are briefly discussed.

Text-fig. 1. Geographical and geological framework. A: map of Egypt with position of Wadi Qena (B); B map of Wadi Qena with position of study area (C); C: geological

map of study area (modified after Conoco, 1987)

GEOLOGICAL SETTING

Wadi Qena is located in the central part of theEgyptian Eastern Desert (Text-fig. 1A, B). The mainpart of Wadi Qena is about 200 km long and runs fromthe southern slopes of the Southern Galala Plateau,near the Gulf of Suez, towards the south southeast. Nearthe town of Qena City it grades into the River Nile val-ley. Wadi Qena is bordered by the uniform, elevated andflat-topped Eocene limestone Plateau in the west andthe serrated, varicoloured basement rocks of the RedSea Mountains range in the east (Text-fig. 1B). WadiQena clearly developed along faults that run parallel tothe Gulf of Suez and the Red Sea Mountains, which arewell exposed at numerous places within the wadi, es-pecially on the eastern side (Bandel et al. 1987). Sed-imentary successions are well exposed on the westernflank of the wadi while Quaternary alluvial fans pre-dominate in the centre (Text-fig. 1C).

The sedimentary rocks of Wadi Qena range in agefrom Palaeozoic to Quaternary (Text-fig. 1C). Most ofthe Palaeozoic rocks are non-marine and well devel-oped in the northern part of the wadi. The distributionof the Palaeozoic sediments was more or less con-trolled by older, pre-Cenomanian structural events(Said 1990; Issawi et al. 1999). The first marine trans-gression took place during the Late Cenomanian. Thisresulted in the deposition of shallow marine, mixed sili-ciclastic and carbonate sediments. During Late Ceno-manian–Turonian times, the southern shoreline of theTethyan seaway was situated immediately south of thestudy area. The marine transgression continued withsome phases of regression during the later parts of theLate Cretaceous and Tertiary, leaving Coniacian–Eocene strata in the study area (see Text-fig. 1C).

SECTIONS

Two sections, located on the western margin in thenorthern part of Wadi Qena (Text-fig. 1C), have beenmeasured bed-by-bed. One section, measured at N 27о

57` 57``, E 32о 30` 58``, is termed the north Wadi Qenasection (Text-fig. 2). The other section, located 25 kmsouth of the north Wadi Qena section, was measured atthe mouth of Wadi El-Burga entering Wadi Qena at N27о 44` 51``, E 32о 33` 55`` and is consequently termedthe Wadi El-Burga section (Text-fig. 2).

The sedimentary succession starts with the Palaeo-zoic Naqus Formation (non-marine, kaolinitic,medium- to coarse-grained sandstones; see Abdallah etal. 1992 and Issawi et al. 1999) which, in places,overlies basement rocks unconformably. Along a trun-

cated palaeosol, the Naqus Formation is overlain bythe Galala Formation. The Galala Formation (Text-fig.2) starts with silty, glauconitic sandstones rich in thetrace fossil Thalassinoides isp. and fragmented mol-lusc shells, indicating shallow marine conditions.Higher up-section, shale beds are common (especiallyin the north), with a few sandy limestone and thin bio-turbated sandstone intercalations. The lower part of theGalala Formation (35 m in the north and 25 m in thesouth) is terminated by a fossiliferous limestone bed,yielding a Late Cenomanian faunal assemblage in-cluding Inoceramus ex gr. pictus Sowerby (Nagm etal. 2011). The following interval consists of gypsifer-ous shale in the north and silty shale with thin sand-stone interbeds in the south. It contains a highly fos-siliferous limestone bed around the ~55-m-level (fromthe base of the Galala Formation) in both sections.This bed is characterized by a typical Late Ceno-manian fauna represented by oysters [Ceratostreonflabellatum (Goldfuss), Curvostrea rouvillei (Co-quand), Ilymatogyra africana (Lamarck)], nautiloids[Angulithes mermeti (Coquand)], ammonites [Neolo-bites vibrayeanus (d’Orbigny)] and echinoids (Hemi-aster cubicus Desor) indicating a fully marine, la-goonal environment (cf. Nagm 2009; Wilmsen andNagm 2012). Marls with calcareous nodules are com-mon in the following part, which ends at a sharp andmore or less irregular surface at 70 m and 64 m re-spectively in the two sections. The upper part of theGalala Formation (50 m in the north and 45 m in thesouth) is characterized by a high carbonate contentand consists of thick limestone beds intercalating withmarl containing calcareous nodules and/or sandy silt-stone (in the south). These sediments are highly fos-siliferous, containing abundant latest Cenomanian andEarly Turonian ammonites (most of the studied spec-imens have been collected from this interval). TheGalala Formation is truncated along a prominent un-conformity at the base of the Umm Omeiyid Forma-tion. This unconformity has also been recognized inWadi Araba and placed in the Lower–Middle Turon-ian boundary interval (Nagm et al. 2010a, b). Thethickness of the Galala Formation is about 120 m inthe North Wadi Qena section and about 112 m in theWadi El-Burga section.

The Umm Omeiyid Formation overlies the GalalaFormation unconformably, attaining a thickness ofabout 50 m in both sections. It consists of unfossilif-erous fissile siltstone with thin, medium-grained, inpart hummocky cross-stratified sandstone intercala-tions. The upper part of the formation is also charac-terized by siliciclastic facies, but additionally containsa few fossiliferous limestone beds and/or marl inter-

65CENOMANIAN – TURONIAN AMMONITES FROM EGyPT


Text-fig. 2. Graphic logs of the North Wadi Qena and Wadi El-Burga sections with stratigraphic distribution of the zonal ammonite markers. The dashed lines represent

inferred ranges of taxa depending on other stratigraphic data. For location see Text-fig. 1C


calations. The calcareous layers yield abundant earlyLate Turonian ammonites [Coilopoceras requieni-anum (d’Orbigny)]. The Umm Omeiyid Formation isoverlain by the Coniacian Taref Sandstone Formationalong a sharp, ferruginous surface.

SySTEMATIC PALAEONTOLOGy

Repositories of specimens: 90 specimens (mostlycomposite internal moulds) form the basis of the study.Illustrated material is kept in the Museum für Miner-alogie und Geologie (MMG) of the SenckenbergNaturhistorische Sammlungen Dresden (SNSD), Ger-many (repository AfK). Additional specimens arehoused in the Department of Geology, Al-Azhar Uni-versity, Egypt (repository AUEN).

Dimensions: All linear dimensions, taken with aVernier Caliper, are given in millimetres. The abbre-viations that are used in the present work are: maxi-mum diameter (D), whorl breadth (Wb), whorl height(Wh), breadth of umbilicus (U). Figures in parenthe-ses are dimensions as a percentage of diameters.

Suture terminology: The suture terminology fol-lows Wedekind (1916) as applied by Kullman andWiedmann (1970) and revised in parts by Korn et al.(2003): E=external lobe, A=adventive lobe (=laterallobe, L, of former nomenclature), U=umbilical lobe,I=internal lobe.

Systematics: The ammonites studied are arrangedsystematically according to Wright et al. (1996). Theterminology used for the description of the taxa fol-lows the glossary in the Treatise on Invertebrate Pale-ontology, Part L, Mollusca 4 (Wright et al., 1996). Inorder to keep this article as short as possible, the syn-onymy lists only contain a few selected referenceswhich are important for the discussion.

Order Ammonoidea von Zittel, 1884Suborder Ammonitina Hyatt, 1889

Superfamily Hoplitoidea Douvillé, 1890? Family Engonoceratidae Hyatt, 1900

Genus Neolobites Fischer, 1882

TyPE SPECIES: Ammonites vibrayeanus d’Orbigny,1841; by original designation.

Neolobites vibrayeanus (d’Orbigny, 1841)(Text-figs 3, 6A)

1841. Ammonites vibrayeanus d’Orbigny, 322, pl. 96, figs1–3.

1981. Neolobites vibrayeanus (d’Orbigny); Kennedy andJuignet, 23, figs 3–4, 6a; text-fig. 5.

1989. Neolobites vibrayeanus (d’Orbigny); Luger andGröschke, 366, pl. 39, fig. 3, text-fig. 5.

2005. Neolobites vibrayeanus (d’Orbigny); Wiese andSchulze, 933, figs 4–9. [with synonymy]

2010a. Neolobites vibrayeanus (d’Orbigny); Nagm et al.,475, figs 4, 5A–F.

MATERIAL: Twenty specimens (AfK 297–301, AUEN1–15) from three different levels in the middle part of theGalala Formation in both sections (Text-fig. 2).

MEASUREMENTS:Specimen D Wb Wh Wb/Wh U

AfK 298 86.4 (100) 20.6 (23.8) 51.0 (59.0) 0.40 8.1 (9.3)

AfK 299 92.9 (100) 20.2 (21.7) 51.7 (55.6) 0.39 8.8 (9.4)

AfK 300 57.3 (100) 16.1 (28.0) 33.7 (58.8) 0.47 4.9 (8.5)

AfK 301 72.5 (100) 18.3 (25.2) 46.9 (64.6) 0.39 4.4 (6.0)

DESCRIPTION: Most of the specimens are fully sep-tate internal moulds with very involute coiling. The

Text-fig. 3. Whorl section and suture line of Neolobites vibrayeanus (d’Orbigny, 1841), Upper Cenomanian of the Galala Formation, specimen AfK 301

small umbilicus, comprising about 6% of the diameter,is shallow, with low and rounded umbilical walls. Theflanks are slightly convex with a narrow and flat ven-ter. The whorl section (Text-fig. 3) is highly com-pressed, elevated, and has a lanceolate shape withgreatest breadth around mid-flank. The suture lines(Text-fig. 3) are poorly preserved and characterized bybroad saddles with somewhat narrower and lanceolatelobes that become shallower towards the umbilicus.The ornamental features are also poorly preserved ex-cept for some delicate clavi at the ventrolateral margin(e.g., Text-fig. 6A4–5).

REMARKS: Despite the imperfect preservation, thespecimens collected from the Galala Formation ofWadi Qena can be assigned to N. vibrayeanus with-out any doubt. Neolobites includes several speciesand many morphological variants, the differences be-tween which have been discussed by Kennedy andJuignet (1981) and Wiese and Schulze (2005). Most,if not all, Neolobites specimens that have beenrecorded from Egypt are representatives of onespecies, i.e., N. vibrayeanus. The morphologicalvariation in the Egyptian material (presence or ab-sence of very weak ribs and ventrolateral clavi) is re-lated to different taphonomic conditions (for moredetails on N. vibrayeanus from Egypt see Nagm etal. 2010a).

OCCURRENCE: Neolobites vibrayeanus is a wide-spread and characteristic early Late Cenomanianspecies, occurring in France, Spain, Portugal, Mo-rocco, Algeria, Tunisia, Egypt, Israel, Lebanon, Syria,Saudi Arabia, Oman, Niger, Bolivia, Colombia, Peruand Venezuela. According to Wiese and Schulze(2005), the species preferred shallow-water environ-ments in subtropical/tropical areas. In the study area,N. vibrayeanus characterizes the lower Upper Ceno-manian Neolobites vibrayeanus Zone, which is (asdefined by Nagm et al. 2010b) equivalent to the Ceno-manian standard ammonite Zone of Calycoceras(Proeucalycoceras) guerangeri (cf. Kennedy 1984;Wright et al. in Wright and Kennedy 1984; Hancock1991).

Superfamily Acanthoceratoidea de Grossouvre, 1894Family Acanthoceratidae de Grossouvre, 1894Subfamily Euomphaloceratinae Cooper, 1978

Genus Euomphaloceras Spath, 1923

TyPE SPECIES: Ammonites euomphalus Sharpe,1855; by monotypy.

Euomphaloceras costatum Cobban, Hook andKennedy, 1989

(Text-figs 4, 6B)

1989. Euomphaloceras costatum Cobban, Hook andKennedy, 37, figs 37, 77S–EE, 78A–H.

2005. Euomphaloceras costatum Cobban, Hook andKennedy; Gale et al.

MATERIAL: Three specimens (AfK 305–307) fromthe upper part of the Galala Formation (97 m above thebase) at the Wadi El-Burga section (Text-fig. 2).


AfK 306 64.8 (100) 32.3 (49.8) 25.7 (39.6) 1.25 23.8 (36.7)

DESCRIPTION: The coiling is evolute, with a largeumbilicus and rounded umbilical walls. The whorlsection (Text-fig. 4) is quadratic, with the greatestbreadth at the umbilical tubercles. Flanks are narrow,flattened and ribbed, with strong umbilical and ven-trolateral tubercles occurring on all main ribs. Theventer is broad and slightly convex, showing a weakridge at the site of the siphuncle, and carries threerows of blunt tubercles (the middle one on the siphonalridge). The suture line is poorly preserved.

REMARKS: The specimens described herein are veryclose to the holotype as described by Cobban et al.(1989). They differ in their less dense ribbing. In spiteof their imperfect preservation, they are assigned to E.costatum without reservation.

OCCURRENCE: Upper Cenomanian of New Mexico(USA), southern England and Brazil. In the study area,the species has been collected from the upper part ofthe Vascoceras cauvini Zone, which is (as defined byNagm et al. 2010b) equivalent to the Neocardiocerasjuddii Zone, in the standard ammonite zonation(Kennedy 1984; Wright et al. in Wright and Kennedy1984; Hancock 1991). This is the first record of E.costatum from Egypt, which confirms the temporal


Text-fig. 4. Whorl section of Euomphaloceras costatum Cobban, Hook and

Kennedy, 1989, Upper Cenomanian of the Galala Formation, specimen AfK 305

equivalence of the V. cauvini Zone to the N. juddiiZone (see below).

Genus Pseudaspidoceras Hyatt, 1903

TyPE SPECIES: Ammonites footeanus Stoliczka,1864; by original designation.

Pseudaspidoceras sp.(Text-figs 5, 6C)

MATERIAL: One specimen (AfK 304) from the upperpart of the Galala Formation (95 m above the base) atthe Wadi El-Burga section (Text-fig. 2).


AfK 304 95.0 (100) 36.9 (38.8) 53.5 (56.3) 0.68 29.3 (30.8)

DESCRIPTION: The coiling is moderately evolute,with a large umbilicus (ca. 30%). The whorl section(Text-fig. 5) is trapezoidal, with the strongest infla-tion at the umbilical shoulders. From the umbilicalshoulders, the flanks converge towards a feebly con-vex venter with narrowly rounded ventrolateralshoulders. The ornamentation consists of distant,rounded and sometimes branched ribs carrying um-bilical bullae. Each rib ends at the ventrolateral shoul-der with a strong tubercle. Ribbing weakens towardsthe outer whorl. The suture line (Text-fig. 5) is wellpreserved and characterized by a broad adventivelobe.

REMARKS: This well preserved specimen has all thediagnostic characters of the genus Pseudaspidoceras.However, it is difficult to identify at the specific levelbecause of the inadequate match of its morphologicalcharacters with those of known species of the genus.In fact, there are only two species that come close tothe present specimen, i.e., P. conciliatum Stoliczka var.ampakabensis Collignon (1965, p. 36, fig. 1669) andP. pseudonodosoides Choffat (1898, p. 65, pl. 16, figs5–8; pl. 22, figs 32–33). However, P. conciliatumStoliczka var. ampakabensis differs from our specimenin a slightly concave venter and highly indented su-tures; furthermore, it is known from the Lower Tur-onian whereas the specimen from Wadi Qena was col-lected from uppermost Cenomanian strata. Instratigraphic position, P. pseudonodosoides matchesour specimen, but it differs morphologically in its de-pressed whorl section with a very broadly roundedventer. The Wadi Qena specimen possibly representsa new species of the genus Pseudaspidoceras, but thisconclusion cannot be based on a single specimen.

OCCURRENCE: The specimen was collected fromthe upper Galala Formation at the Wadi El-Burga sec-tion, 2 m above the level with Vascoceras cauvini and2 m below the level with Euomphaloceras costatum.Therefore, its stratigraphic position is equivalent to theupper Upper Cenomanian Neocardioceras juddii stan-dard ammonite Zone (sensu Wright et al. in Wright andKennedy 1984).

Family Vascoceratidae Douvillé, 1912Genus Vascoceras Choffat, 1898


Text-fig. 5. Suture line and whorl section of Pseudaspidoceras sp., Upper Cenomanian of the Galala Formation, specimen AfK 304


Text-fig. 6. A – Neolobites vibrayeanus (d’Orbigny, 1841) (A1, A2 - lateral and ventral views of AfK 298, ×1; A3 - lateral view of AfK 300, ×1; A4, A5 - lateral

and ventral views of AfK 301, ×1). B – Euomphaloceras costatum Cobban, Hook and Kennedy, 1989 (B1, B2 - lateral and ventral views of AfK 305, ×1).

C – Pseudaspidoceras sp. (C1, C2 - lateral and ventral views of AfK 304, ×1). All specimens from the Upper Cenomanian of the Galala Formation


TyPE SPECIES: Vascoceras gamai Choffat, 1898;by the subsequent designation of Diener, 1925.

Vascoceras cauvini Chudeau, 1909(Text-figs 7, 9A)

1909. Vascoceras cauvini Chudeau, 67, pl. 1, figs 1a, 2a; pl.2, figs 3, 5; pl. 3, figs 1b, 2b, 4.

1969. Paravascoceras cauvini (Chudeau); Freund and Raab,20, pl. 3, figs 1–3; text-fig. 5a, b.

1989. Vascoceras cauvini Chudeau; Luger and Gröschke,374, pl. 40, figs 3, 6, 8, 9; pl. 41, figs 1–4; pl. 42, fig.1; text-figs 6g,h, 8c.

2010a. Vascoceras cauvini Chudeau; Nagm et al., 483, figs9A–B, 10B.

MATERIAL: Two specimens (AfK 302, 303) from theupper part of the Galala Formation (93 m above thebase) at the Wadi El-Burga section (Text-fig. 2).


AfK 302 96.3 (100) 41.3 (42.8) 36.1 (37.4) 1.14 33.4 (34.6)

AfK 303 98.0 (100) 34.3 (35.0) 41.5 (42.3) 0.82 37.1 (37.8)

DESCRIPTION: Both specimens are fragmentary in-ternal moulds with relatively evolute coiling as well asa deep and wide umbilicus (ca. 35%). The umbilicalshoulders are feebly rounded with vertical walls.Flanks are short and convex with a broadly rounded tonearly flat venter (in the latter case, faintly defined ven-trolateral shoulders are present). The whorl section iscompressed to slightly depressed (Text-fig. 7, speci-men AfK 302). The ornamentation is not preserved inspecimen AfK 302 (Text-fig. 9A3–4), while veryweak, shallow ribs, especially towards the venter, oc-cur in specimen AfK 303 (Text-fig. 9A1–2, a fragmentof a body chamber). The suture line, characterized bya deep external lobe and a wide, deep adventive lobe,is clearly visible in specimen AfK 302 (see Text-fig. 7).

REMARKS: In Vascoceras cauvini two forms areknown (see Chudeau 1909); i.e. the smooth form(e.g. Luger and Gröschke 1989; Zaborski 1996; Alyet al. 2008; Barroso-Barcenilla and Goy 2010; Nagmet al. 2010a); and the ribbed form (e.g., Freund andRaab 1969; Zaborski 1990, 1996; Meister et al.1992). Two different forms also occur in the studyarea: specimen AfK 302 is a typical representative ofthe smooth form (Text-fig. 9A3–4) while specimenAfK 303 (Text-fig. 9A1–2) shares characters of boththe smooth and ribbed forms in terms of dimensionsand suture line. On the other hand, it has its own char-acters, i.e., a flat venter and the absence of the pro-nounced ornamentation typical of the ribbed form(very weak, shallow ribs occur only at the venter).Therefore, this specimen is considered here as a tran-sitional morph between the smooth and ribbed formof V. cauvini.

Vascoceras pioti (Peron and Fourtau, 1904; inFourtau, 1904) likewise has a flat venter, but itsumbilicus is very small and the suture is different:the lobes of V. cauvini are deeper and wider thanthose of V. pioti, especially the A-lobe (Text-fig. 7).In addition, the suture line of V. cauvini has threesaddles, while that of V. pioti has four (see Freundand Raab 1969, p. 30, text-fig. 6f). The sutures of V.cauvini from Wadi Qena (Text-fig. 7) are identicalto sutures in the type material as illustrated byChudeau (1909). Furthermore, V. pioti is from ahigher stratigraphic level, coming from the LowerTuronian.

OCCURRENCE: Vascoceras cauvini is widely knownfrom the Upper Cenomanian of Nigeria, Niger, Angola,Morocco, Algeria, Egypt, Israel, Spain and Peru. In thestudy area, Vascoceras cauvini characterizes the up-permost Cenomanian Vascoceras cauvini Zone, whichis (cf. Nagm et al. 2010b) equivalent to the Neocar-dioceras juddii Zone in the standard ammonite zona-tion of Kennedy (1984), Wright et al. in Wright andKennedy (1984) and Hancock (1991).

Text-fig. 7. Whorl section and suture line of Vascoceras cauvini Chudeau, 1905, Upper Cenomanian of the Galala Formation, specimen AfK 302


Vascoceras globosum globosum (Reyment, 1954)(Text-figs 8, 9B)

1954. Pachyvascoceras globosum Reyment, 259, pl. 3, fig.3; pl. 5, fig. 4; text-figs 3e, 7.

1957. Vascoceras globosum globosum (Reyment); Barber,21, pl. 7, figs 1a, b; pl. 9, figs 4a, b; pl. 28, figs 1, 2.

2005. Vascoceras globosum globosum (Reyment); Gale etal., 182, fig. 9.

MATERIAL: A single specimen (AfK 308) from theupper part of the Galala Formation (102 m above thebase) at the Wadi El-Burga section (Text-fig. 2).


AfK 308 80.3 (100) 69.2 (86.1) 41.5 (51.6) 1.66 10.1 (12.5)

DESCRIPTION: The specimen is a completely smooth,globose, very involute phragmocone with a maximumpreserved diameter of 80.3 mm. The umbilicus is small(12.5% of diameter) and very deep, with a narrowlyrounded umbilical shoulder. The flanks are short andstrongly convex, grading without ventrolateral shoul-ders into a broadly rounded venter. The whorl section(Text-fig. 8) is depressed and globose, with a whorl-breadth to whorl-height ratio of 1.66. The suture line iswell preserved and illustrated in Text-fig. 8.

REMARKS: Barber (1957) erected five subspecies ofVascoceras globosum based on variations in the degreeof inflation. Of those subspecies, V. globosum globosumis the most inflated. The specimen from Wadi Qena isstrongly depressed and globose in shape, with a smalland deep umbilicus. These characters are diagnostic ofVascoceras globosum globosum as described and il-lustrated by Barber (1957) and Zaborski (1996).

OCCURRENCE: Uppermost Cenomanian of Nigeriaand Lower Turonian of Brazil. In the study area, it has

been recorded from the Lower Turonian, i.e., the lowerpart of the Choffaticeras spp. Zone. This zone is (as de-fined by Nagm et al. 2010b) equivalent to the upperpart of the standard ammonite Zone of Watinocerascoloradoense (Kennedy 1984; Wright et al. in Wrightand Kennedy 1984) or Watinoceras spp. (Hancock1991). This is the first record of Vascoceras globosumglobosum from Egypt.

Vascoceras kossmati Choffat, 1898(Text-figs 9C, 10, 13A)

1898. Vascoceras kossmati Choffat, 63, pl. 13, figs 8, 9; pl.14, figs 1, 2; pl. 21, figs 26, 27.

1992. Paravascoceras kossmati (Choffat); Thomel, 225, pl.119, figs 2, 3.

2010. Vascoceras kossmati Choffat; Barroso-Barcenilla and Goy,219, pl. 5, figs F–H; text-fig. 6B. [with synonymy].

MATERIAL: Four specimens (AfK 309–312) from theupper part of the Galala Formation (102 m above thebase) at the Wadi El-Burga section (Text-fig. 2).


AfK 309 65.5 (100) 48.0 (73.2) 30.2 (46.1) 1.58 13.2 (20.1)

AfK 310 52.5 (100) 39.6 (75.4) 27.4 (52.1) 1.44 9.0 (17.1)

AfK 311 59.9 (100) 46.4 (77.4) 33.0 (55.0) 1.40 7.8 (13.0

AfK 312 66.4 (100) 45.6 (68.6) 35.2 (53.0) 1.29 9.1 (13.7)

DESCRIPTION: All specimens are smooth internalmoulds characterized by strongly involute coiling. Theumbilicus is narrow and deep, showing vertical wallsand sharp margins. The flanks are strongly convex,grading into a broadly rounded venter. Some speci-mens (AfK 311, 312) have wide and flat venters thatare clearly separated from the flanks by ventrolateralshoulders (Text-fig. 13A2). The whorl section (Text-fig. 10) is depressed, with an incipiently triangular

Text-fig. 8. Whorl section and suture line of Vascoceras globosum globosum (Reyment, 1954), Lower Turonian of the Galala Formation, specimen AfK 308


Text-fig. 9. A – Vascoceras cauvini Chudeau, 1909 (A1, A2 - ventral and lateral views of AfK 303, ×1; A3, A4 - lateral and ventral views of AfK 302, ×1; Upper

Cenomanian of the Galala Formation). B – Vascoceras globosum globosum (Reyment, 1954) (B1, B2 - lateral and ventral views of AfK 308, ×1; Lower Turonian

of the Galala Formation). C – Vascoceras kossmati Choffat, 1898 (C1, C2 - lateral and apertural views of AfK 309, ×1; Lower Turonian of the Galala Formation)


shape. The suture line (Text-fig. 10) is simple andcharacterized by a wide and deep adventive lobe.

REMARKS: Barroso-Barcenilla and Goy (2010) dis-cussed the similarities between Vascoceras kossmatiand Vascoceras hartti (Hyatt). They regarded the sub-spherical shape, the small umbilical width and the de-pressed whorl section as the key characters of V. koss-mati. The material from Wadi Qena is very close to theSpanish material, and we thus assign it to Choffat’sspecies without hesitation.

OCCURRENCE: Lower Turonian of Portugal, Spainand France (see Barroso-Barcenilla and Goy 2010).Eck (1914) noted the species from Egypt without il-lustration. Therefore, this is the first properly docu-mented record from Egypt. In the study area, it oc-curs in the Lower Turonian, in the lower part of theChoffaticeras spp. Zone, the equivalent of the upperpart of the standard ammonite Zone of Watinocerascoloradoense (Kennedy 1984; Wright et al. inWright and Kennedy 1984) or Watinoceras spp.(Hancock 1991) (see Nagm et al. 2010b for furtherdetails).

Genus Neoptychites Kossmat, 1895

TyPE SPECIES: Ammonites telinga Stoliczka, 1865;by the subsequent designation of Solger 1904 (= Am-monites cephalotus Courtiller 1860).

Neoptychites cephalotus (Courtiller, 1860)(Text-figs 11, 13B)

1860. Ammonites cephalotus Courtiller, 248, pl. 2, figs 1–4.1907. Neoptychites cephalotus (Courtiller); Pervinquière,

393, pl. 27, figs 1–4, text-fig. 152.

1994. Neoptychites cephalotus (Courtiller); Chancellor etal., 70, pl. 16, figs 1–9; pl. 17, figs 1–5; pl. 18, figs1–3; pl. 26, figs 2–4. [with synonymy]

2008. Neoptychites cephalotus (Courtiller); Kennedy et al.,159, pl. 3, figs 6–8; pl. 6, figs 1–5.

2010a. Neoptychites cephalotus (Courtiller); Nagm et al.,489, figs 11G–H, 12A–B, 13A–B.

MATERIAL: Three specimens (AfK 313, AUEN 16–17) from the upper part of the Galala Formation at thenorth Wadi Qena section, co-occurring with Choffat-iceras segne (Text-fig. 2).


AfK 313 63.3 (100) 42.9 (67.7) 32.9 (51.9) 1.30 5.1 (8.0)

DESCRIPTION: All three specimens are characterizedby very involute coiling with a small and shallow um-bilicus (U ~8% of the diameter). The greatest whorlbreadth is situated shortly above the umbilical shoul-der. Whorls are generally depressed, with a subtrigo-nal section (Text-fig. 11). The flanks are strongly con-vex, converging to a rounded venter. Ornamentationand suture line are not seen due to the poor preserva-tion.

Text-fig. 10. Whorl section and suture line of Vascoceras kossmati Choffat, 1898, Lower Turonian of the Galala Formation, specimen AfK 309

Text-fig. 11. Whorl section of Neoptychites cephalotus (Courtiller, 1860),

Lower Turonian of the Galala Formation, specimen AfK 313

REMARKS: Large collections of Neoptychitescephalotus have been described from Tunisia (Pervin-quière 1907; Chancellor et al. 1994), France (Kennedyand Wright 1979), and New Mexico, USA (Cobban andHook 1983). The last authors demonstrated a widerange of intraspecific variation and dimorphism in thespecies. This variation has also been documented inEgyptian material by Nagm et al. (2010a). The presentmaterial falls within the range of variation described inthe literature; it is close to the Tunisian material (Chan-cellor et al. 1994) and material described recently fromMorocco (Kennedy et al. 2008).

OCCURRENCE: Known from France, northern Spain,Morocco, Algeria, Tunisia, Egypt, Israel, Syria, Jordan,Cameroon, Madagascar, southern India, Colorado,New Mexico (USA), northern Mexico, Trinidad,Venezuela, Colombia, Brazil, Niger and Nigeria. In thestudy area, this species co-occurs with Choffaticerassegne and Thomasites rollandi which are dominant inthe Lower Turonian of Egypt (Nagm et al. 2010a, b).It ranges into the basal Middle Turonian Collignon-iceras woollgari Zone (Kennedy et al. 2008).

Family Pseudotissotiidae Hyatt, 1903Subfamily Pseudotissotiinae Hyatt, 1903

Genus Thomasites Pervinquière, 1907

TyPE SPECIES: Pachydiscus rollandi Thomas andPeron, 1889; by subsequent designation of Diener,1925.

Thomasites gongilensis (Woods, 1911)(Text-figs 12, 13C)

1911. Vascoceras gongilensis Woods, 282, pl. 21, fig. 7; pl.22, fig. 1.

1981. Thomasites gongilensis (Woods); Wright andKennedy, 100, pl. 24, fig. 1; pl. 25, fig. 1; including va-rieties tectiformis Barber 1957, and lautus Barber1957. [with full synonymy]

1989. Thomasites gongilensis (Woods); Meister, 38, pl. 16,figs 3–5; pl. 17, figs 1–6; pl. 18, figs 1–3; pl. 19, figs1–5; pl. 20, figs 1–5; pl. 21, figs 1–3.

2003. Thomasites gongilensis (Woods); Kennedy et al., 12,pl. 5, figs 1–3.

MATERIAL: One specimen (AfK 314) from the upperpart of the Galala Formation at the north Wadi Qenasection, co-occuring with Choffaticeras segne (Text-fig. 2).


AfK 314 81.8 (100) 42.3 (51.7) 43.5 (53.1) 0.97 19.3 (23.5)

DESCRIPTION: The specimen has a preserved diam-eter of ~82 mm and a moderately wide, deep umbili-cus with broadly rounded umbilical shoulders. Thewhorl section (Text-fig. 12) is only slightly compressed(Wb/Wh = 0.97), with the greatest width at the um-bilical shoulders. The flanks are convex, ending atrounded ventrolateral shoulders. The venter is fairlybroad and flat. The suture line (Text-fig. 12) shows nar-row lobes and broad, weakly incised saddles. Orna-mental features are not preserved.

REMARKS: Wright and Kennedy (1981) revisedthe genus Thomasites and discussed the relationshipof the present species to other species referred to thegenus. West African faunas of T. gongilensis (e.g.,


Text-fig. 12. Whorl section and suture line of Thomasites gongilensis (Woods, 1911), Lower Turonian of the Galala Formation, specimen AfK 314


Text-fig. 13. A – Vascoceras kossmati Choffat, 1898 (A1, A2 - lateral and ventral views of AfK 311, ×1). B – Neoptychites cephalotus (Courtiller, 1860) (B1, B2

- apertural and ventral views of AfK 313, ×1). C – Thomasites gongilensis (Woods, 1911) (C1, C2 - apertural and lateral views of AfK 314, ×1). D – Thomasites

rollandi (Thomas and Peron, 1889) (D1, D2 - apertural and lateral views of AfK 316, ×1). All specimens from the Lower Turonian of the Galala Formation


Barber 1957) have been compared with Tunisianmaterial of T. rollandi by Chancellor et al. (1994).They noted that T. gongilensis rarely shows coarseumbilical tubercles (in contrast to juveniles of T.rollandi), and generally has a less inflated, less tri-angular whorl section as well as a simpler suture line.The present specimen of T. gongilensis differs fromT. rollandi in a wider umbilicus and a simple suture.The Sinai specimen of T. gongilensis of El-Hedeny(2002, fig. 4d, e) most probably represents Vasco-ceras cauvini.

OCCURRENCE: Thomasites gongilensis (Woods1911) is common in the Lower Turonian of WestAfrica. Kennedy et al. (2003) recorded it from theUpper Cenomanian of France. In the study area, ithas been recorded from the Lower Turonian Chof-faticeras spp. Zone, which is (see Nagm et al.2010b) equivalent to the upper part of the standardammonite Zone of Watinoceras coloradoense(Kennedy 1984; Wright et al. in Wright andKennedy 1984) or Watinoceras spp. (Hancock1991). This is the first unequivocal record of T.gongilensis from Egypt.

Thomasites rollandi (Thomas and Peron, 1889)(Text-figs 13D, 14, 16A)

1889. Pachydiscus Rollandi Thomas and Peron, 25, pl. 17,figs 1–3.

1994. Thomasites rollandi (Thomas and Peron); Chancel-lor et al., 75, pl. 19, figs 1–2; pl. 20, figs 1–12; pl. 21,figs 1–9; pl. 22, figs 1–6; pl. 23, figs 1–9; text-fig.14a–f. [with synonymy]

2010a. Thomasites rollandi (Thomas and Peron); Nagm etal., 493, figs 13E–G, 14A–D, 15A–D.

MATERIAL: Seven specimens (AfK 315, 316; AUEN18–22) from the upper part of the Galala Formation at theWadi El-Burga section, co-occuring with Choffaticerassegne (Text-fig. 2).


AfK 315 88.7 (100) 47.1 (53.1) 50.2 (56.5) 0.93 10.3 (11.6)

AfK 316 88.9 (100) 49.6 (55.7) 51.1 (57.4) 0.97 9.3 (10.4)

DESCRIPTION: The incompletely preserved internalmoulds are smooth and characterized by involute coil-ing (U ~10% of diameter). The umbilical wall is high,subvertical and ends at narrowly rounded umbilicalshoulders. The greatest whorl breadth is at the umbil-ical shoulder. The flanks are generally weakly convexand converge strongly to a narrowly rounded venter.The whorl section is compressed to slightly depressedand shows a trigonal (Text-fig. 14) to ovoidal shape(specimen AfK 316; Text-fig. 13D). The suture line(Text-fig. 14) is characterized by four saddles, the onebetween the external and adventive lobes being high,broad and strongly incised.

REMARKS: Thomasites rollandi has been extensivelyrevised by Chancellor et al. (1994). They clarified therange of its intraspecific variation and discussed its re-lationship to other species of the genus. Into syn-onymy of Th. rollandi they put numerous varietiesand species of Pervinquière (1907) and Douvillé(1928). The preservation of the specimens studied isnot good but they clearly belong to T. rollandi, espe-cially specimen AfK 316 (Text-fig. 13D), which isvery close to the holotype as re-figured in Chancelloret al. (1994, pl. 19).

OCCURRENCE: The species is well known from

Text-fig. 14. Whorl section and suture line of Thomasites rollandi (Thomas and Peron, 1889), Lower Turonian of the Galala Formation, specimen AfK 315

the Lower Turonian of Morocco, Tunisia, Algeria,Egypt, Israel, Jordan, Syria, Lebanon, France andSpain. It may also occur in England, Colombia andTadzhikistan (Chancellor et al. 1994). In the studyarea, T. rollandi co-occurs with Lower TuronianChoffaticeras segne, the range of which equates tothe upper part of the Watinoceras coloradoenseZone of Kennedy (1984) and Wright et al. in Wrightand Kennedy (1984) (see Nagm et al. 2010b for de-tails).

Genus Pseudotissotia Peron, 1897

TyPE SPECIES: Ammonites galliennei d’Orbigny,1850; by subsequent designation of Pervinquière,1907.

Pseudotissotia nigeriensis (Woods, 1911)(Text-figs 15, 16B)

1911. Hoplitoides nigeriensis Woods, 284, pl. 23, fig. 3; pl.24, fig. 15; text-fig. 1.

1989. »Pseudotissotia« nigeriensis (Woods); Meister, 44,pl. 21, figs 4–6; pl. 22, figs 1–4; pl. 23, figs 1–5; pl.24, fig. 1; pl. 25, figs 1–7; pl. 26, figs 1–3; pl. 27, figs1–3.

2005. Pseudotissotia nigeriensis (Woods); Gale et al., 186,figs 10D–E, 12A–B, E, 13A–B.

MATERIAL: Eleven specimens (AfK 328–338) fromthe uppermost part of the Galala Formation, in bothsections studied (~110 m above the base of the for-mation; Text-fig. 2).


AfK 328 68.5 (100) 19.7 (28.7) 37.5 (54.7) 0.36 6.0 (8.7)

AfK 329 91.1 (100) 33.4 (36.6) 50.3 (55.2) 0.66 11.7 (12.8)

AfK 330 84.0 (100) 20.1 (23.9) 43.0 (51.1) 0.46 9.8 (11.6)

AfK 331 85.8 (100) 27.3 (31.1) 47.0 (54.7) 0.58 7.7 (9.0)

AfK 332 78.5 (100) 23.4 (29.8) 42.3 (53.8) 0.55 8.2 (10.4)

DESCRIPTION: All specimens are smooth, fully sep-tate internal moulds with a maximum preserved diam-eter of 91 mm (specimen AfK 329, Text-fig. 16B4–6).Coiling is involute (U ~10%) with a moderately deepumbilicus and feebly rounded umbilical walls. Theflanks are high, subparallel, with convex inner andconverging outer parts. Two prominent keels at theventrolateral shoulders mark the contact with the ven-ter, which is characterized by a siphonal keel, resultingin the distinctive tricarinate appearance of the ventralarea (Text-fig. 15). The three keels have the same rel-ative height in all specimens. The whorl section (Text-fig. 15) is compressed, with the maximum breadth di-rectly above the umbilical shoulder. The suture line(Text-fig. 15) is characterized by very broad saddles andnarrow lobes; the adventive lobe is deeply incised.

REMARKS: The flat, tricarinate venter, the com-pressed whorl section and a suture line with broadsaddles and narrow lobes (deeply incised adventivelobe) are characteristic of Pseudotissotia nigeriensis(e.g., Meister 1989). Based on these characters, thespecimens from the upper Galala Formation of WadiQena are unequivocally attributed to Woods’ speciesand represent its first proven occurrence in Egypt.Our specimens closely match the Nigerian material asdescribed by Woods (1911) and Meister (1989).

EMAD NAGM AND MARKUS WILMSEN78

Text-fig. 15. Whorl section and suture line of Pseudotissotia nigeriensis (Woods, 1911), Lower Turonian of the Galala Formation, specimen AfK 328

CENOMANIAN – TURONIAN AMMONITES FROM EGyPT79

Text-fig. 16. A – Thomasites rollandi (Thomas and Peron, 1889) (A1, A2 –lateral and apertural views of AfK 315, ×1). B – Pseudotissotia nigeriensis (Woods, 1911) (B1,

B2 - lateral and apertural views of AfK 328, ×1; B3 - lateral view of AfK 334, ×1; B4, B5, B6 - apertural, lateral and ventral views of AfK 329, ×1). All specimens from

the Lower Turonian of the Galala Formation

OCCURRENCE: The species is known from theLower Turonian of Nigeria, Algeria, Israel, northernMexico and Brazil. In the study area, it occurs in theLower Turonian, above the mid-Lower Turonian Chof-faticeras spp. Zone. The P. nigeriensis horizon is thusconsidered to be equivalent to the lower part of thestandard ammonite Zone of Mammites nodosoides (cf.Kennedy 1984; Wright et al. in Wright and Kennedy1984; Hancock 1991), since the last appearance ofChoffaticeras in Egypt is very close to the interpolatedfirst occurrence of Mammites nodosoides (Nagm et al.2010b; see Text-fig. 22 and discussion below).

Genus Choffaticeras Hyatt, 1903

TyPE SPECIES: Pseudotissotia meslei Peron, 1897,by original designation.

Subgenus Choffaticeras Hyatt, 1903

Choffaticeras (Choffaticeras) securiforme (Eck, 1909)(Text-fig. 17)

1909. Tissotia securiformis Eck, 187, figs 9–13.1969. Choffaticeras securiforme (Eck); Freund and Raab,

60, pl. 9, fig. 1; text-fig. 11i–k.1996. Choffaticeras gr. securiforme (Eck); Meister and Ab-

dallah, 15, pl. 9, fig. 1; text-fig. 6g.2009. Choffaticeras (Choffaticeras) securiforme (Eck);

Nagm, 91, text-figs 5.19, 5.20A, 5.23H, I.

MATERIAL: Two fragmentary specimens (AUEN23–24) from the upper part of the Galala Formation

(102 m above the base of the formation) at the NorthWadi Qena section (Text-fig. 2).

DESCRIPTION: The specimens are large and charac-terized by a wide and fairly deep umbilicus, with ver-tical walls and a sharp umbilical shoulder. The flanksconverge to a narrow venter with a strong median, ob-tuse keel. The whorl section (Text-fig. 17) is com-pressed, lanceolate in shape and with the maximumbreadth at the umbilical shoulder. Ornament and sutureline are not preserved.

REMARKS: Freund and Raab (1969) described a largecollection (49 specimens) of Choffaticeras securiformefrom Israel, and discussed its relationship to C. (Leon-iceras) luciae (Pervinquière) and C. (L.) massipianum(Pervinquière). Although the specimens studied are verypoorly preserved, their whorl section (Text-fig. 17) andthe narrow venter, with a conspicuous median obtusekeel, allow their safe assignment to C. securiforme.

OCCURRENCE: The species is known from theLower Turonian of Israel, Egypt and Tunisia. In thestudy area, it comes from the middle Lower TuronianChoffaticeras spp. Zone, which is (as defined by Nagmet al. 2010b), equivalent to the upper part of the stan-dard ammonite Zone of Watinoceras coloradoense(Kennedy 1984; Wright et al. in Wright and Kennedy1984) or Watinoceras spp. (Hancock 1991).

Choffaticeras (Choffaticeras) segne (Solger, 1903)(Text-figs 18, 19)

1903. Pseudotissotia segnis Solger, 77, pl. 4, figs 1, 2; text-figs 16–21.

1904. Schloenbachia quaasi Peron, 255, pl. 1, figs 1–3.2007. Choffaticeras (Choffaticeras) quaasi (Peron); Bar-

roso-Barcenilla and Goy, 464, figs 4(4,5), 5(1).2007. Choffaticeras (Choffaticeras) segne (Solger); Bar-

roso-Barcenilla and Goy, 468, figs 5(5), 6(1–3).2008. Choffaticeras (Choffaticeras) segne (Solger); Ken-

nedy et al., 162, fig. 9; pl. 3, figs 4, 5; pl. 7, figs 1, 2;pl. 8, figs 1–6.

2009. Choffaticeras (Choffaticeras) quaasi (Peron); Nagm,88, text-figs 5.18A, 5.23C.

2009. Choffaticeras (Choffaticeras) segne (Solger); Nagm,94, text-figs 5.17B, 5.18C, 5.23E–G.

MATERIAL: Twenty-seven specimens (AfK 317–327, AUEN 25–33) from the upper part of the GalalaFormation (103–104 m above the base of the forma-tion) in both studied sections (cf. Text-fig. 2).


Text-fig. 17. Whorl section of Choffaticeras (Choffaticeras) securiforme

(Eck, 1909), Lower Turonian of the Galala Formation, specimen AUEN 23

from north Wadi Qena Section


AfK 317 101.3 (100) 36.8 (36.3) 50.2 (49.5) 0.73 13.0 (12.8)

AfK 318 107.4 (100) 31.8 (29.6) 47.5 (44.2) 0.66 17.4 (16.2)

AfK 319 114.1 (100) 36.9 (31.9) 67.0 (56.2) 0.55 23.1 (20.2)

AfK 320 81.8 (100) 22.5 (27.5) 42.9 (52.4) 0.52 10.0 (12.2)

AfK 321 57.9 (100) 20.0 (34.5) 33.7 (58.2) 0.59 3.6 (6.2)

AfK 322 45.5 (100) 15.4 (33.8) 25.9 (56.9) 0.59 5.8 (12.7)

AfK 323 43.8 (100) 12.9 (29.4) 23.8 (54.3) 0.54 5.4 (12.3)

DESCRIPTION: Both small and large specimens arecharacterized by moderately involute coiling. The um-bilicus has outward-inclined walls and rounded um-bilical shoulders. The flanks are high and generallyconvex, especially on the inner parts. Early whorls arecommonly ornamented by simple ribs and very weakventrolateral tubercles that disappear during ontogeny.The venter is tricarinate in juvenile and sharp in adultstages. The whorl section is compressed, with an oxy-cone shape (Text-fig. 18) and the greatest breadth onthe inner part of the flanks. The suture line is well pre-served in all specimens and shows some variability(Text-fig. 18). However, the sutures are generally char-acterized by broad and high saddles with a deep andbroad adventive lobe.

REMARKS: Barroso-Barcenilla and Goy (2007) de-scribed several species of Choffaticeras from Spainand discussed their relationship. The Lower Turonianof Egypt yielded abundant Choffaticeras (cf. Nagm2009; Nagm et al. 2010a, b); however, only twospecies are represented in Wadi Qena, i.e., Choffat-iceras (Choffaticeras) securiforme and Choffaticeras(C.) segne (and eight species are known in contempo-raneous strata of Wadi Araba in the northern part of theEastern Desert; Nagm 2009; Wilmsen and Nagm, inprep.). The well-preserved and large population of C.(C.) segne from the Eastern Desert confirms the viewof Kennedy et al. (2008) that C. (C.) quaasi is a jun-ior synonym of C. (C.) segne (Wilmsen and Nagm, inprep.).

OCCURRENCE: The species is known from theLower Turonian of Tunisia, Israel, Syria, Jordan,France and Spain. In the study area, it characterizes themiddle Lower Turonian Choffaticeras spp. Zone,which is (as defined by Nagm et al. 2010b) equivalentto the upper part of the standard Watinoceras colora-doense Zone (Kennedy 1984; Wright et al. in Wrightand Kennedy 1984) or the Watinoceras spp. Zone(Hancock 1991).


Text-fig. 18. Whorl section and suture line of Choffaticeras (Choffaticeras) segne (Solger, 1903), Lower Turonian of the Galala Formation, A, B - specimen AfK 321;

C - specimen AfK 317; D - specimen AfK 323


Text-fig. 19. Choffaticeras (Choffaticeras) segne (Solger, 1903) from the Lower Turonian of the Galala Formation. A1, A2 - lateral and apertural views of AfK 319, ×1;

A3, A4 - apertural and lateral views of AfK 317 , ×1; A5, A6 - apertural and lateral views of AfK 325, ×1

Family Coilopoceratidae Hyatt, 1903Genus Coilopoceras Hyatt, 1903

TyPE SPECIES: Coilopoceras colleti Hyatt, 1903, byoriginal designation.

Coilopoceras requienianum (d’Orbigny, 1841)(Text-figs 20, 21)

1841. Ammonites requienianum d’Orbigny, 315, pl. 93,figs 1–4.

1984. Coilopoceras requienianum (d’Orbigny); Kennedyand Wright, 282, pls. 35, 36, text-figs 1–5. [withsynonymy]

1989. Coilopoceras requienianum (d’Orbigny); Luger and Gröschke, 388, pl. 46, text-figs 6a, e, 11, 12, 13a–c.

2010a. Coilopoceras requienianum (d’Orbigny); Nagm etal., 495, figs 16A–C, 17, 18A–C.

MATERIAL: Fifteen specimens (AfK 339–341,AUEN 34–45) from the upper part of the UmmOmeiyid Formation in both sections studied (Text-fig. 2).


AfK 339 93.9 (100) 31.0 (33.0) 61.7 (65.7) 0.50 5.8 (6.1)

AfK 340 95.6 (100) 36.0 (37.6) 63.1 (66.0) 0.57 5.9 (6.1)

AfK 341 100.7 (100) 36.4 (36.1) 59.5 (59.0) 0.61 4.9 (4.8)

DESCRIPTION: All specimens are characterized byvery involute coiling. The umbilicus is small (U ~5–6%) and narrow, with low and rounded shoulders. Theflanks are convex and converge towards a pointedventer. The whorl section (Text-fig. 20) is compressed,with a lanceolate shape and the maximum breadth onthe inner flanks. Most specimens are smooth but someof the small specimens have a few weak ribs on the in-ner whorls. The suture line (Text-fig. 20) shows abroad and shallow external lobe, and a very broad,open and bifid adventive lobe.

REMARKS: Cobban and Hook (1980) demonstratedthat many of the described species of the genusCoilopoceras are probably synonyms, often beingbased on small differences in the suture lines. Theirview is supported by the re-description of the type ma-terial of C. requienianum by Kennedy and Wright(1984), who suggested the existence of sexual dimor-phism in C. requienianum shown by smooth and ribbedforms. The specimens from Wadi Qena closely corre-spond to the type material. However, some are more in-flated and the smooth morphotype predominates.

OCCURRENCE: Upper Turonian of Madagascar,Egypt, North and South America, France and Spain. Inthe study area, C. requienianum characterizes the lowerUpper Turonian zone of Coilopoceras requienianum,which is (as defined by Nagm et al. 2010b) equivalentto the lower part of the standard ammonite Zone ofSubprionocyclus neptuni (Hancock 1991).


Text-fig. 20. Suture line and whorl section of Coilopoceras requienianum (d’Orbigny, 1841), Upper Turonian of the Umm Omeiyid Formation, specimen AfK 339


Text-fig. 21. Coilopoceras requienianum (d’Orbigny, 1841) from the Upper Turonian of the Umm Omeiyid Formation. A1, A2 - lateral and apertural views of AfK 339, ×1;

A3, A4 - apertural and lateral views of AfK 341, ×1

BIOSTRATIGRAPHICAL AND PALAEOBIOGEO-GRAPHICAL REMARKS

The first marine transgression in the study area tookplace during the Late Cenomanian, documented by thedeposition of the shallow marine Galala Formationabove the Palaeozoic Naqus Formation, with a promi-nent sequence boundary (major non-sequence) at thebase of the Galala Formation. The early Late Ceno-manian is known as a time of major eustatic sea-levelrise (e.g., Wilmsen 2003; Wilmsen et al. 2005). Thelower part of the Galala Formation is characterized byglauconitic, silty sediments, rich in oysters distinctive ofUpper Cenomanian successions elsewhere in Egypt(Nagm 2009). Approximately 35 m above the base ofthe formation, the sediments have yielded the earlyLate Cenomanian ammonite Neolobites vibrayeanus(see Wiese and Schulze 2005 for details of the species).It defines the oldest ammonite zone in the study area(Text-fig. 2), equivalent to the northwest European stan-dard ammonite Zone of Calycoceras (Proeucalyco-ceras) guerangeri (Kennedy 1984; Wright et al. inWright and Kennedy 1984; Hancock 1991) and theTethyan standard ammonite Zone of Calycoceras nav-iculare (Text-fig. 22; see Nagm et al. 2010b for full de-tails). The beds of the Galala Formation below the firstoccurrence of N. vibrayeanus are included in the N. vi-brayeanus Zone (cf. Nagm 2009; Nagm et al. 2010b).N. vibrayeanus ranges for about 35 m above its first oc-currence, defining the top of the zone (Text-fig. 2). It isfollowed by an ammonite-barren 20–25 m interval richin the oyster Costagyra olisiponensis. Based on the lat-est Cenomanian age of the overlying strata, the barreninterval is regarded here as equivalent to the middle Up-per Cenomanian Metoicoceras geslinianum Zone ofKennedy (1984), Wright et al. in Wright and Kennedy(1984) and Hancock (1991; see Text-fig. 22).

Above the barren interval, ammonites are com-mon again. The lowest eight metres are characterizedby high carbonate contents and yield, in the lowerpart, Vascoceras cauvini. This is the index of theeponymous uppermost Cenomanian ammonite zone inEgypt, largely equivalent to the Neocardioceras juddiiZone of Kennedy (1984), Wright et al. in Wright andKennedy (1984) and Hancock (1991), in the Ceno-manian standard ammonite zonation (Nagm et al.2010b). Euomphaloceras costatum has been recordedfrom the upper part of the Vascoceras cauvini Zone, forthe first time from Egypt. This species co-occurs withNeocardioceras juddii in the United States (Cobban etal. 1989). In the study area, the upper limit of this zoneis considered to mark the Cenomanian–Turonian (C–T) boundary (cf. Nagm et al. 2010b).

Two metres above the inferred C–T boundary (ashort distance below the 100-m-level above the base ofthe Galala Formation in both sections; Text-fig. 2),Early Turonian vascoceratids (V. globosum globosumand V. kossmati) become abundant in high-carbonatesediments. The absence of Vascoceras proprium andPseudaspidoceras flexuosum suggests a stratigraphicgap at this level (see Text-fig. 22), comparable to thesituation in some sections of Wadi Araba in the north-ern part of the Eastern Desert (Nagm et al. 2010b).Two horizons with representatives of the genus Chof-faticeras have been recognized directly above the low-ermost Turonian vascoceratid interval (Text-fig. 2): thelower one with Choffaticeras (Choffaticeras) securi-forme; and the upper one with C. (C.) segne. The ~7 mthick Choffaticeras interval characterizes the middleLower Turonian (Nagm et al. 2010b), equivalent to theupper part of the standard ammonite Zone ofWatinoceras devonense (Kennedy 1984; Hancock1991; see Text-fig. 22). This part is followed by thehighly fossiliferous uppermost interval of the GalalaFormation, yielding typical Nigerian ammonites, suchas Pseudotissotia nigeriensis (Text-fig. 2), dated as lateEarly Turonian (Meister 1989). In agreement with thisobservation, Nagm et al. (2010b) proposed that the lastappearance of the genus Choffaticeras approximatesthe first appearance of the index of the standard upperLower Turonian Mammites nodosoides ammoniteZone of Kennedy (1984) and Hancock (1991). There-fore, the P. nigeriensis Zone (or horizon) seems to be


Text-fig. 22. Ammonite biostratigraphy of the Upper Cenomanian and Lower

Turonian. Vertically hatched fields represent stratigraphic gaps. See text for

further explanations

equivalent to the lower part of the M. nodosoidesZone. The upper boundary of this zone is marked bya prominent unconformity in the Lower–Middle Tur-onian boundary interval (Text-fig. 2). The same un-conformity has been recognized in the sections ofWadi Araba in the northern part of the Eastern Desert;Nagm et al. 2010b), and it also has its equivalent inshallow shelf settings of the northern Tethys margin(see Niebuhr et al. 2011). This unconformity also rep-resents the contact of the Galala and Umm Omeiyidformations. The lower part of Umm Omeiyid Forma-tion is dominated by siliciclastics and is barren ofmacrofauna. In the upper part of the formation, fos-siliferous strata yielded the early Late Turonian am-monite Coilopoceras requienianum, the zone of whichis (as defined by Nagm et al. 2010b) equivalent to thelower part of the Upper Turonian standard ammoniteZone of Subprionocyclus neptuni (Geinitz) (Kennedy1984; Hancock 1991; see Text-figs 2, 22). This obser-vation suggests that the unfossiliferous part of thelower Umm Omeiyid Formation corresponds, in abroad sense, to the Middle Turonian. Interestingly,also in the sections of Wadi Araba, Middle Turonianstrata are barren of macrofossils (Nagm 2009; Nagmet al. 2010a, b).

The early Late Cretaceous eustatic transgression,with a significant sea-level rise that peaked during theLate Cenomanian and Early Turonian, has been welldocumented (Hancock and Kauffman 1979; Hardenbolet al. 1998; Sharland et al. 2001; Wilmsen 2003; Wilm-sen et al. 2005, 2010). Rising sea level may be astraightforward explanation for the wide geographicdistribution of many Cenomanian–Turonian am-monites (Wiedmann 1988; Lehmann and Herbig 2009;Nagm et al. 2010a). The palaeontological record doc-uments ammonite geographic expansion during theLate Cenomanian, expressed by the presence of a sin-gle large Tethyan province, followed by biogeographicdifferentiation during the Early Turonian, expressed byBoreal, Vascoceratid and Puzosiid provinces (Wied-mann 1988; Lehmann and Herbig 2009). The palaeo-biogeographic affinities of the studied fauna corre-spond to this pattern (see Nagm et al. 2010a for datafrom Wadi Araba). The early and mid-Late Cenoman-ian ammonite fauna is biogeographically not very di-agnostic. The latest Cenomanian incipient provincial-ism is documented by the presence of the typicallyNorth and West African Vascoceras cauvini. The EarlyTuronian assemblage, studied herein, belongs to theTethyan Vascoceratid Province, with strong influencesof Nigerian faunas (Vascoceras globosum globosum,Thomasites gongilensis, Pseudotissotia nigeriensis),proving the faunal connection to Central and West

Africa via the Trans-Saharan Seaway (see Collignonand Lefranc 1974; Meister et al. 1992). The palaeo-biogeographic affinity to the Boreal Realm is muchless noticeable in the study area, with only 50% of theammonites of Wadi Qena known also from that Realm.It is noteworthy that, in the contemporaneous succes-sions of the northern part of the Eastern Desert (WadiAraba, ca. 130 km to the north), 85% of the Early Tur-onian ammonoids identified are also known from theBoreal Realm (Nagm et al. 2010a). This observationmay indicate relatively steep north–south environ-mental gradients.

Acknowledgements

We greatly acknowledge constructive reviews by F.Barroso-Barcenilla (Madrid), C. Meister (Geneva) and ananonymous referee. Furthermore, we thank W.J. Kennedy(Oxford) for the discussion of taxonomic problems as wellas I. Walaszczyk (Warszawa) and C.J. Wood (Minehead) fortheir editorial work. R. Winkler and M. Röthel (SenckenbergDresden) did the photographic and collection work atSNSD. The financial support of DAAD for the stay of ENin Germany is greatly acknowledged. This paper is a con-tribution to German Research Foundation (DFG) project WI1743/6-1.

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